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Who Needs Emotions? The Brain Meets the Robot

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366 conclusions<br />

from <strong>the</strong> primary taste cortex and <strong>the</strong> primary olfactory (pyriform) cortex;<br />

from <strong>the</strong> amygdala; and from midbrain dopamine neurons. As Rolls documents,<br />

damage to <strong>the</strong> caudal orbitofrontal cortex produces emotional changes, which<br />

include <strong>the</strong> tendency to respond when responses are inappropriate (i.e., <strong>the</strong><br />

tendency of monkeys not to withhold responses to nonrewarded stimuli). Rolls<br />

sees orbitofrontal neurons as part of a mechanism which evaluates whe<strong>the</strong>r a<br />

reward is expected and generates a mismatch (evident as a firing of <strong>the</strong> nonreward<br />

neurons) if <strong>the</strong> reward is not obtained when it is expected.<br />

As Fellous and LeDoux note, decision-making ability in emotional situations<br />

is also impaired in humans with damage to <strong>the</strong> medial prefrontal cortex<br />

and abnormalities in <strong>the</strong> prefrontal cortex may predispose people to<br />

develop fear and anxiety disorders. <strong>The</strong>y suggest that <strong>the</strong> medial prefrontal<br />

cortex allows cognitive information processing in <strong>the</strong> prefrontal cortex to<br />

regulate emotional processing by <strong>the</strong> amygdala, while emotional processing<br />

by <strong>the</strong> amygdala may influence <strong>the</strong> decision-making and o<strong>the</strong>r cognitive<br />

functions of <strong>the</strong> prefrontal cortex. <strong>The</strong>y <strong>the</strong>n suggest that <strong>the</strong> prefrontal–<br />

amygdala interactions may be involved in <strong>the</strong> conscious feelings of fear.<br />

However, this neat division between <strong>the</strong> cognitive cortex and emotional<br />

amygdala strikes me as too glib—both because not all parts of <strong>the</strong> cortex<br />

give rise to conscious feelings and because human emotions seem to be inextricably<br />

bound up with “cortical subtleties.”<br />

Neuromodulation<br />

We have now seen something of <strong>the</strong> crucial roles of <strong>the</strong> hypothalamus,<br />

amygdala, and orbitofrontal cortex in <strong>the</strong> motivational system. In a similar<br />

vein, Fellous (1999) reviewed <strong>the</strong> involvement of <strong>the</strong>se three areas in emotion<br />

and argued that <strong>the</strong> neural basis for emotion involves both computations<br />

in <strong>the</strong>se structures and <strong>the</strong>ir neuromodulation. It is thus a useful feature<br />

of <strong>the</strong> present volume that Kelley’s analysis of motivation and emotion<br />

emphasizes three widely distributed chemical signaling systems and <strong>the</strong>ir<br />

related functions across different phyla. Following are some key points from<br />

her richly detailed survey.<br />

We first discuss dopamine, reward, and plasticity. In mammals, dopamine<br />

is proposed to play a major role in motor activation, appetitive motivation,<br />

reward processing, and cellular plasticity and may well play a major<br />

role in emotion. In <strong>the</strong> mammalian brain, dopamine is contained in specific<br />

pathways, which have <strong>the</strong>ir origins in <strong>the</strong> substantia nigra pars compacta and<br />

<strong>the</strong> ventral tegmental area of <strong>the</strong> midbrain and ascend to innervate widespread<br />

regions of striatal, limbic, and cortical regions such as <strong>the</strong> striatum,<br />

prefrontal cortex, amygdala, and o<strong>the</strong>r forebrain regions. Studies in <strong>the</strong>

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